We took a short break from the Alienist books two weeks ago to recognise our author’s birthday with the first half of a special investigative history blog examining the truth behind the legendary panthers of Davon Wood who feature in The Legend of Broken, Mr. Carr’s 2012 epic set in Northern Germany during the Dark Ages. We discovered in Part One that the impressive proportions of “Davon panthers”–that is, nine-foot bodies from nose to tail base and weights of over 500lbs (226kg)–ruled out the possibility of the great cats having been relatives of either the European (or Eurasian) jaguar (Panthera gombaszoegensis) or the ancestors of modern African lions (Panthera leo), and instead favoured the hypothesis that Davon panthers would more likely have been relatives of the equally impressive European (or Eurasian) cave lion (Panthera spelaea spelaea)1, a Pleistocene felid species capable of reaching weights of up to 617lbs (280kg).

Having established that the lengths and weights of the Davon panther fall into the same ranges as those of the European cave lion, we open Part Two with the more intriguing question of lifestyle: do the hunting styles, prey choices, and environmental preferences of Davon panthers also support the European cave lion hypothesis? We will be reviewing fascinating research into mitochondrial DNA and bone collagen isotopic signatures to answer this question, as well as investigating possible extinction hypotheses for the European cave lion and the implications these have for the likelihood of Davon panthers having really survived through to the Dark Ages.

Lone cats, not lone wolves.

We are told in The Legend of Broken that Davon panthers were solitary hunters, a characteristic of their species that might initially appear to be at odds with the hypothesis that these legendary cats were relatives of the European cave lion. Until recently, the European cave lion was assumed to hunt cooperatively in a large pride structure like the modern lion. Palaeolithic cave paintings such as the panel found in the Chauvet Cave in Ardèche, France, which depicts a large number of cave lions in close proximity to one of their proposed prey species, the bison, has been taken as evidence in support of this assumption (Bocherens et al., 2011). Moreover, the assumed diet of the cave lion, which includes large ungulate prey such as horse, bison, reindeer, and woolly rhinoceros, along with juvenile members of megaherbivore species such as mammoth, has been modelled on the diet of the modern lion, a species that relies on cooperative hunting methods to bring down the large adult ungulates (zebra and antelope) that form the foundation of their diet, and will even prey on the modern equivalents of megaherbivores–juvenile weaned elephants–when prides are large enough (i.e., up to 30 members) to support the collective hunting required to feasibly target such large prey (Bocherens et al., 2011). However, two mitochondrial DNA studies (Burger et al., 2004; Barnett et al., 2009) and two bone collagen isotopic signature studies (Bocherens et al., 2011; Yeakel et al., 2013) have recently called into question the assumption that European cave lions shared behavioural traits such as collective hunting with the modern lion.

During the Pleistocene, felids that resemble the modern lion covered territories from Africa and Europe, through Asia, and into North America. Mitochondrial DNA research carried out on fossil specimens now clearly demonstrates that the European cave lion, the American lion, and the modern lion should be viewed as three distinct species. Specifically, Burger et al. (2004) and Barnett et al. (2009) have both shown that although the American lion (Panthera atrox, see left) and the European cave lion (Panthera spelaea) shared common ancestors approximately 200,000 years ago, both species were “genetically isolated” from the ancestors of modern lions and did not contribute mitochondrial DNA to the modern lion population. This is supported by the additional interesting finding that there were several points of cross-contact between the three species at various stages of the Pleistocene–one area of cross-contact for American lions and European cave lions was located between the Yukon/Alaska and Southern Canada, while another area of extended cross-contact for the ancestors of modern lions and European cave lions was located the Near East–and yet no inter-breeding appears to have taken place, indicating that the American lion, the European cave lion, and the ancestors of modern lions were, indeed, three distinct felid species (Barnett et al., 2009). In addition to shedding further light on felid taxonomy during the Pleistocene, this important finding also brings into question the assumption that European cave lions and American lions shared the collective hunting behaviour of modern lions, a unique trait that is not observed in any other modern Panthera species.

An independent line of enquiry utilising bone collagen isotopic signatures has recently provided more direct evidence for the hunting behaviour of European cave lions and competing predator species throughout Europe during the Pleistocene. Because the ratios of stable nitrogen and carbon isotopes for prey species are recorded in the tissues of predators, Pleistocene predator-prey networks have been able to be reconstructed using bone collagen isotopic signatures to a degree of accuracy that has even allowed the proportional contributions of specific prey species in the diets of different predators to be determined (Yeakel et al., 2013). The first comprehensive study of European cave lion prey choices was conducted by Bocherens et al. (2011) using the fossilised remains of various predator species and their potential prey species from 25 sites around Belgium, Germany, France, and Switzerland that were dated as falling into one of two time periods: before the Last Glacial Maximum (pre-LGM: i.e., approx. 24,000 to 40,000 years ago), and after the Last Glacial Maximum (post-LGM: i.e., approx. 12,000 to 14,000 years ago). Interestingly, the cave lions showed a wide scattering of isotopic values, suggesting that different individuals were preying on species with contrasting isotopic signatures, a finding that the authors argue strongly supports the hypothesis that cave lions were solitary hunters with individual prey preferences rather than hunting collectively in prides (Bocherens et al., 2011). | Continue reading →

In recognition of our author’s birthday today, I’m taking a short break from the Alienist books for a fun diversion (for me, and hopefully for our author, too): an investigative history blog about the legendary panthers found roaming the forests of Northern Germany in his 2012 novel, The Legend of Broken. For those visitors of 17th Street who haven’t read the book, you may be wondering why a big cat is featured on the cover. If this is the case, prepare to be enlightened! For those of you who have read the book, while you will already be familiar with the legendary panthers of Davon Wood, perhaps you are less clear on what species the great cats are most likely to correspond to: Mr. Carr presented the European (or Eurasian) jaguar (Panthera gombaszoegensis) and the European (or Eurasian) cave lion (Panthera spelaea spelaea1) as candidate species2, but refrained from providing an argument in favour of one over the other. In the following post, we will review the evidence before us from the latest research into these ancient big cat species–including two studies published earlier this year–to put forward such an argument, and hopefully illuminate a little more of the true history of these great predators of ancient Europe.

Happy Birthday, Mr. Carr!

Who were the legendary panthers of Davon Wood?

Within The Legend of Broken we are transported back in time to the year 750 A.D., to the heart of the time period we know today as the Dark Ages that spanned the fourth or fifth century A.D. to the eleventh century A.D. The walled city of Broken, located on the peak of the Harz Mountain Range in Northern Germany, is surrounded by rugged terrain, with the Thuringian Forest (known in the novel as “Davon Wood”) bordering the region to the south. It is here that the exiles of Broken, known as the Bane, have learned to live in harmony with a variety of predators, including the most fearsome of them all–the rarely seen, solitary big cats known as “Davon panthers”.

Wolves in Davon Wood grow to extraordinary sizes, and are more than a match for any three Bane—even these three. Keera, however, shakes her head slowly, and answers: “A panther.” Veloc’s face, too, fills with apprehension, while Heldo-Bah’s shows childlike panic. The solitary, silent Davon panthers–which can reach lengths of twelve feet, and weights of many hundreds of pounds–are the largest and most efficient killers known, each as lethal as a pack of wolves and, like all cats, nearly impossible to detect before they strike. They are particularly fond of the caves and rocks near the Cat’s Paw.

When we finally do meet one of these legendary panthers, we can understand the awe with which the Bane regard these magnificent creatures. Described as reaching proportions of over nine feet from nose to tail base–twelve feet with tail included–with coat colours ranging from darkest gold to the palest gold/sand adorned with rich dark spots and stripes, presumably on the underside of the body and legs, it is not difficult to imagine why the cats achieved legendary status with their human cohabitants. Solitary hunters, they seem to have learned to survive on the deer, boar, and wild horses that roam the forest, or even to venture close to human habitation to poach shag cattle when times are tough.

However, instead of simply accepting the existence of the impressive “Davon panthers” as the Bane do, we twenty-first century readers are compelled to ask a very twenty-first century question upon meeting these remarkable creatures (or at least, this twenty-first century woman is): What are these great cats? Our author tells us in the endnotes of the novel that, “The Legendary ‘European panther’ is far more than a myth,” and goes on to cite two potential candidate species: the smaller European jaguar (Panthera gombaszoegensis), and the larger European cave lion (Panthera spelaea spelaea), both of which could be found over an enormous geographical range that included Northern Germany during the Pleistocene, an epoch that ended approximately 11,000 years ago. See the map below for a overview of the known geographic range of the European cave lion in and around Northern Germany (two key locations for The Legend of Broken–the city of Broken itself and Davon Wood–are also marked), including radiocarbon dates of known cave lion finds. Perhaps the reason our author refrained from providing an argument for one species over the other was due to the the fact that both the jaguar and the cave lion had their strengths and weaknesses as candidate species. Nevertheless, on the basis of very recent research, I propose that the strengths of one candidate species now outweighs the strengths of the other. So, let’s revisit the mystery and review the evidence as it currently stands.

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Broken (Brocken)

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The location of the city of Broken in The Legend of Broken.

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Forest where the legendary panthers roam in The Legend of Broken. One is said to live in a cave in a remote corner of the forest.

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In the case of the Davon panther, size matters.

The first and most important evidence to review in order to solve the mystery of the Davon panther relates to the sizes of the candidate species under consideration. After all, repeated descriptions of Davon panthers as having nine foot bodies from nose to tail base with weights of over 500lbs (226kg) simply can’t be disregarded, and as the European jaguar and cave lion were in entirely different size brackets, the proportions of the cats must be considered the first and most important differentiator between the two species. Perhaps the most compelling evidence in relation to size comes from a recent paper by Marciszak (2014) who conducted a comprehensive overview of size variability among European jaguars in the late Middle Pleistocene in relation to contemporary European Panthera species including the European cave lion and the European leopard on the basis of remains found for all three species collected from Biśnik Cave, Poland, in 2010.

On the basis of tooth size, previous research has established approximate body mass ranges for European jaguars of 330lbs to 418lbs (150-190kg) for males, and 220lbs to 286lbs (100-130kg) for females. Despite the fact that the jaguar from layer 19d of Biśnik Cave was evaluated to be a large, male specimen at the top of the size spectrum with an estimated body mass of 396lbs (180kg), the specimen was still strikingly smaller than contemporary cave lions found in layers 19b-d of Biśnik Cave (see the figure drawn to scale from Marciszak, 2014, below). Importantly, after reviewing specimens from almost all sites where fossil material for the European jaguar has been found, no size increase was observed for the species over its evolution, indicating that there is no reason to suppose the European jaguar would have almost doubled its maximum body mass by the period in which The Legend of Broken was set, assuming they had not died out by that period. On the other hand, the estimated weights for European cave lions do fall well within the range proposed for Davon panthers with a maximum estimated weight of up to 793lbs (360kg) for Panthera spelaea fossilis and 617lbs (280kg) for Panthera spelaea spelaea (Guzvica, 1998), although changes in size for the species throughout its evolution have recently been shown to be more complex than previously theorised.

It is now generally accepted that the earliest cave lions in Europe (Panthera spelaea fossilis), which first appeared during the Middle Pleistocene, were gradually replaced by the more evolutionarily advanced Panthera spelaea spelaea during the Late Pleistocene. By once again performing comparative analyses of specimens from the palaeontologically important Biśnik Cave in Poland, Marciszak and Stefaniak (2010) observed a clear evolutionary progression from fossilis in the Middle Pleistocene, through intermediate forms that shared features with both fossilis and spelaea in the late Middle Pleistocene, to the cave lion’s final spelaea form in the Late Pleistocene. Although this evolutionary progression involved changes to cranial and dental morphology, as well as an overall decrease in body size, a more recent review of the evolution of Panthera spelaea from the fossilis to spelaea subspecies by Marciszak, Schouwenburg, and Darga (2014) includes data from a new specimen found in the San River in Poland which has shown that the evolutionary decrease in size may not have been as definitive as the 2010 study suggested.

Although both Radiocarbon and Uranium-Thorium dating methods failed on the San River specimen, the advanced cranial and dental morphology of the specimen is consistent with a male Late Pleistocene Panthera spelaea spelaea, most probably from the first half of the last glacial. Nonetheless, no intact male spelaea skull from the last glacial has exceeded 400mm (1.31 foot) in length, while the San River specimen was 451mm (1.48 foot) in length (see the image adapted from Marciszak et al., 2014, to the left), suggesting a body size in line with earlier fossilis forms. Consequently, the authors suggest that either the San River specimen is from an earlier period than the cranial and dental morphology indicate, which they consider to be unlikely, or it may simply be evidence that cave lions with impressive dimensions did continue to survive into the Late Pleistocene, even if such specimens were rare. Taken in combination with the fact that the number of fossilis and spelaea specimens is currently so limited that it is not possible to perform a statistically reliable analysis of European cave lion size at the present time (Marciszak et al., 2014), this is a particularly important finding to take into consideration when assessing candidate species for Davon panthers as the decreasing size for the European cave lion might otherwise be considered problematic for the hypothesis that the panthers were living descendants of Panthera spelaea spelaea.

First, one would presume that over the several thousand years between the Late Pleistocene when Panthera spelaea spelaea appears to have died out and the time period in which The Legend of Broken is set, the size of any remaining cave lions (assuming it was possible for any to have survived) would have continued to decrease, not maintain their former megafauna proportions. Indeed, by the end of the cave lion reign in Europe in the first part of the last glacial, “dwarf” cave lions with dimensions similar to medium-sized modern African lions were appearing among the European cave lion population. The finding of a fossilis-sized cave lion from a much later time period than had previously been thought possible for the Late Pleistocene Panthera spelaea spelaea therefore helps the argument that separate lineages of cave lions may have continued to exist in Europe (see also Guzvica, 1998, for further evidence of separated lineages in European cave lion populations), which further helps the case that Davon panthers may have retained their impressive megafauna dimensions over several thousand years.

Second, it is also important to note that, like the European jaguar, the European cave lion was strongly sexually dimorphic with male skulls typically exceeding female skulls by an average of 3.93 inches (10cm) in length. Given the fact that it was a female Davon panther that was said to exceed 500lbs (226kg) in The Legend of Broken, in order for the immature male panther we meet early in the novel to grow to the correspondingly large size the female’s weight would indicate, the only possible way this could have happened is if the original megafauna proportions had been retained. Incidentally, this point also rules out the possibility that Davon panthers were relatives of the ancestors of modern African lions. Although isolated finds indicate that modern lions colonised areas of South-Western Europe as late as the Iron Age (Stuart & Lister, 2011), there is no evidence that they would have reached the proportions required for a Davon panther, nor that any separate line of modern lion would have evolved to increase in size rather than decrease in size.

To Be Continued

Although the size requirements for Davon panthers have now ruled out the possibility of the great cats having evolved from European jaguars or the ancestors of modern African lions, we are faced with the question of the panthers’ hunting and living habits. Within The Legend of Broken we are told that Davon panthers were solitary hunters rather than hunting collaboratively in prides. Does this correspond to the hypothesised hunting pattern of European cave lions? Perhaps surprisingly, recent mitochondrial DNA and bone collagen isotopic signature findings suggest that it does! We will overview this fascinating research, as well as investigating possible extinction hypotheses and the implications these have for the likelihood of Davon panther existence, in Part Two of The Legendary Panthers of Davon Wood series over the coming weeks. | Continue reading →